Bradley J. Sinclair

Episodic radiations in the fly tree of life

Flies are one of four superradiations of insects (along with beetles, wasps, and moths) that account for the majority of animal life on Earth. Diptera includes species known for their ubiquity (Musca domestica house fly), their role as pests (Anopheles gambiae malaria mosquito), and their value as model organisms across the biological sciences (Drosophila melanogaster). A resolved phylogeny for flies provides a framework for genomic, developmental, and evolutionary studies by facilitating comparisons across model organisms, yet recent research has suggested that fly relationships have been obscured by multiple episodes of rapid diversification. We provide a phylogenomic estimate of fly relationships based on molecules and morphology from 149 of 157 families, including 30 kb from 14 nuclear loci and complete mitochondrial genomes combined with 371 morphological characters. Multiple analyses show support for traditional groups (Brachycera, Cyclorrhapha, and Schizophora) and corroborate contentious findings, such as the anomalous Deuterophlebiidae as the sister group to all remaining Diptera. Our findings reveal that the closest relatives of the Drosophilidae are highly modified parasites (including the wingless Braulidae) of bees and other insects. Furthermore, we use micro-RNAs to resolve a node with implications for the evolution of embryonic development in Diptera. We demonstrate that flies experienced three episodes of rapid radiation—lower Diptera (220 Ma), lower Brachycera (180 Ma), and Schizophora (65 Ma)—and a number of life history transitions to hematophagy, phytophagy, and parasitism in the history of fly evolution over 260 million y.

The phylogenetic relationships among infraorders and superfamilies of Diptera based on morphological evidence

Members of the megadiverse insect order Diptera (flies) have successfully colonized all continents and nearly all habitats. There are more than 154 000 described fly species, representing 10–12% of animal species. Elucidating the phylogenetic relationships of such a large component of global biodiversity is challenging, but significant advances have been made in the last few decades. Since Hennig first discussed the monophyly of major groupings, Diptera has attracted much study, but most researchers have used non‐numerical qualitative methods to assess morphological data. More recently, quantitative phylogenetic methods have been used on both morphological and molecular data. All previous quantitative morphological studies addressed narrower phylogenetic problems, often below the suborder or infraorder level. Here we present the first numerical analysis of phylogenetic relationships of the entire order using a comprehensive morphological character matrix. We scored 371 external and internal morphological characters from larvae, pupae and adults for 42 species, representing all infraorders selected from 42 families. Almost all characters were obtained from previous studies but required revision for this ordinal‐level study, with homology assessed beyond their original formulation and across all infraorders. We found significant support for many major clades (including the Diptera, Culicomorpha, Bibionomorpha, Brachycera, Eremoneura, Cyclorrhapha, Schizophora, Calyptratae and Oestroidea) and we summarize the character evidence for these groups. We found low levels of support for relationships between the infraorders of lower Diptera, lower Brachycera and major lineages of lower Cyclorrhapha, and this is consistent with findings from molecular studies. These poorly supported areas of the tree may be due to periods of rapid radiation that left few synapomorphies in surviving lineages.

Global diversity of dipteran families (Insecta Diptera) in freshwater (excluding Simulidae, Culicidae, Chironomidae, Tipulidae and Tabanidae)

Today’s knowledge of worldwide species diversity of 19 families of aquatic Diptera in Continental Waters is presented. Nevertheless, we have to face for certain in most groups a restricted knowledge about distribution, ecology and systematic, particularly in the tropical environments. At the same time we realize a dramatically decline or even lack of specialists being able, having the time or the opportunity to extend or even secure the present information. The respective families with approximate numbers of aquatic species are: Blephariceridae (308), Deuterophlebiidae (14), Nyphomyiidae (7), Psychodidae (∼2.000), Scatopsidae (∼5), Tanyderidae (41), Ptychopteridae (69), Dixidae (173), Corethrellidae (97), Chaoboridae (∼50), Thaumaleidae (∼170), Ceratopogonidae (∼6.000), Stratiomyidae (∼43), Empididae (∼660), Lonchopteridae (2), Syrphidae (∼1.080), Sciomyzidae (∼190), Ephydridae (∼1.500), Muscidae (∼870). Numbers of aquatic species will surely increase with increased ecological and taxonomical efforts.

Discovered in our backyard: a new genus and species of a new family from the Rocky Mountains of North America (Diptera, Tabanomorpha)

Abstract.  A new genus of the Tabanomorpha, Oreoleptis,gen.n., assigned to the monotypic family Oreoleptidae, fam.n., is described from the Rocky Mountains of North America. The male, female, larva and pupa of Oreoleptis torrenticola, sp.n. are described and illustrated. Adults were reared from larvae collected from torrential streams and rivers, and from pupae collected from riverbanks. No adults have been collected yet in the field. The larvae have two pairs of very long, ventrolateral, crocheted prolegs on abdominal segments 2–7, and a short, dorsal pair on segments 6–7. The larval head and mouthparts resemble those of athericids and tabanids. The mandibular hook has an internal canal, the basal mandibular sclerite is compressed with both condyles coming together and articulating on the tentorial phragma; the mandibular brush is located on a vertical rod; and the salivary pump is greatly enlarged. The adult male genitalia have aedeagal tines similar to athericids, tabanids and Bolbomyia Loew, the hypandrium is fused with the gonocoxites, the epandrium is subrectangular, lying flat on the gonocoxites, and tergite 10 is present (a mixture of advanced and primitive features). The endoaedeagal process is reduced (as in athericids), and the gonocoxal apodemes are long and slender, an advanced condition shared by athericids and tabanids. The female has a long, extrusible postabdomen and ovipositor, two‐segmented cerci, and the basal cercal segment has a prominent posteroventral lobe, typical of rhagionids and pelecorhynchids. Cladistic analysis assigns the genus Oreoleptis to sister group status of the Athericidae + Tabanidae. However, wing venation, simple, unmodified female abdomen, undivided first tergite, and two‐segmented female cercus, excludes this taxon from the Athericidae and Tabanidae. The relationships of the new family are discussed and the phylogeny of the higher Tabanomorpha reassessed.

Male terminalia of Diptera (Insecta): a review of evolutionary trends, homology and phylogenetic implications

The male terminalia character system in Diptera is reviewed. The phylogenetic implications of apomorphic changes are traced on published cladograms. New synapomorphies include: anteroventral parameral apodeme for the Tipulomorpha; parameral sheath encompassing desclerotized aedeagus for Neodiptera (exclusive of Axymyiidae); endoaedeagus for Xylophagomorpha + Tabanomorpha. Apystomyiidae are classified as the sister group to the Eremoneura based on four synapomorphies (lateral ejaculatory processes absent, subepandrial sclerite extending from base of hypoproct to phallus, bacilliform sclerites extending to tips of the epandrium and surstyli functionally developed, but not articulated) and lack of eremoneuran synapomorphies (i.e., loss of gonostyli, presence of postgonites and phallic plate). The Diptera sperm pump with a functional ejaculatory apodeme is a possible autapomorphy of Diptera, exclusive of Nymphomyiidae and Deuterophlebiidae. Internal details of the male terminalia of Sylvicola and Mycetobia (Anisopodidae), Hilarimorpha (Hilarimorphidae) and Apystomyia (Apystomyiidae) are newly illustrated and homologies of the aedeagus, paramere and sperm pump of the Tipuloidea are clarified.

Status of Rhagoletis (Diptera: Tephritidae) Pests in the NAPPO Countries

ABSTRACT The North American Plant Protection Organization (NAPPO) is an organization comprising plant protection regulatory officials of the three signatory countries: the United States, Canada, and México. NAPPO develops Regional Standards for Phytosanitary Measures (RSPMs) as well as discussion papers on important issues related to plant protection. The Fruit Panel of NAPPO organized a Technical Assistance Group (TAG) to develop a discussion paper on the status of pest species of Rhagoletis (Diptera: Tephritidae) within the NAPPO countries. As regulations are developed to prevent the spread of these pests both within and outside of the NAPPO countries, it was important to understand the pest status of these species. This article reviews the basic biology, host range, distribution, potential to spread, management, and regulatory status of pest species of Rhagoletis within the NAPPO countries.

A new species of the seepage midge genus Trichothaumalea Edwards from Japan (Diptera: Thaumaleidae)

The adult and pupa of Trichothaumalea japonica sp. n. are described from Japan. This widespread relict genus also includes three species from western [T. pluvialis (Dyar & Shannon), T. pilosa (Garrett)] and eastern (T. elakalensis Sinclair) North America. T. japonica + T. elakalensis are most closely related and together are the sister-group to the western Nearctic species group. Examples of Diptera with similar biogeographic patterns are listed. In addition, male genitalic homologies in this genus are re-examined.

An update of the blow flies (Diptera: Calliphoridae) of the Galápagos Islands, and first record of Chrysomya rufifacies (Macquart) from mainland Ecuador

Seven species of Calliphoridae are reported from the Galápagos Islands, Ecuador: Lucilia pionia (Walker), L. setosa (James), L. deceptor (Curran), L. eximia (Wiedemann), Cochliomyia macellaria (Fabricius), Chrysomya albiceps (Wiedemann), and Chrysomya megacephala (Fabricius). Lucilia eximia is newly recorded from the islands. Lucilia sp. near pionia is recorded from the island of Española. The distribution and collection records of these species are discussed and listed, and a key to their identification is provided. Chrysomya rufifacies (Macquart) is reported for the first time from mainland Ecuador and the identification of this species is outlined.

Alavesia Waters and Arillo–a Cretaceous-era genus discovered extant on the Brandberg Massif, Namibia (Diptera: Atelestidae)

Two new extant species of the fossil genus Alavesia Waters & Arillo (A. brandbergensissp.n. and A. daurasp.n.) are described from the Brandberg Massif, Namibia. The genus is reassigned to the family Atelestidae (subfamily Atelestinae) on the basis of male terminalia (elongate gonocoxal apodemes, ventral apodemes, subapical surstylus), female terminalia (absence of tergum 10) and a three‐articled stylus. This is the first record of Atelestidae from the Afrotropical Region. A genus‐level phylogeny of the Atelestidae is resolved, in which Alavesia is the sister genus to the remaining genera of the Atelestinae. The biogeographic history of the genus is briefly discussed, as is the significance of this and other relict Diptera on the Brandberg.

Revision of the Nearctic Species of Heleodromia (Diptera: Empidoidea: Brachystomatidae)

Abstract The Nearctic species of the brachystomatid genus Heleodromia Haliday are revised. The following six species are recognized: Heleodromia boreoalpina Saigusa, H. chillcotti Sinclair sp. nov., H. cranehollowensis Cumming and Coovert sp. nov., H. irwini Wagner, H. pullata (Melander), and H. woodi Brooks sp. nov. The following new Palearctic distribution records are also reported: H. boreoalpina (Russian Far East), H. irwini (Germany, Russian Far East), and H. immaculata Haliday (Russian Far East). A key to adults, illustrations of male terminalia, and known distributions are included.